The Influence of Environmental,Biotic and Spatial Factors on Diatom Metacommunity Structure in Swedish Headwater Streams

Stream assemblages are structured by a combination of local (environmental filtering and biotic interactions) and regional factors (e.g., dispersal related processes). The relative importance of environmental and spatial (i.e., regional) factors structuring stream assemblages has been frequently assessed in previous large-scale studies, but biotic predictors (potentially reflecting local biotic interactions) have rarely been included. Diatoms may be useful for studying the effect of trophic interactions on community structure since: (1) a majority of experimental studies shows significant grazing effects on diatom species composition, and (2) assemblages can be divided into guilds that have different susceptibility to grazing. We used a dataset from boreal headwater streams in south-central Sweden (covering a spatial extent of ∼14000 km²), which included information about diatom taxonomic composition, abundance of invertebrate grazers (biotic factor), environmental (physicochemical) and spatial factors (obtained through spatial eigenfunction analyses). We assessed the relative importance of environmental, biotic, and spatial factors structuring diatom assemblages, and performed separate analyses on different diatom guilds. Our results showed that the diatom assemblages were mainly structured by environmental factors. However, unique spatial and biological gradients, specific to different guilds and unrelated to each other, were also evident. We conclude that biological predictors, in combination with environmental and spatial variables, can reveal a more complete picture of the local vs. regional control of species assemblages in lotic environments. Biotic factors should therefore not be overlooked in applied research since they can capture additional local control and therefore increase accuracy and performance of predictive models. The inclusion of biotic predictors did, however, not significantly influence the unique fraction explained by spatial factors, which suggests low bias in previous assessments of unique regional control of stream assemblages.     

A comparative analysis reveals weak relationships between ecological factors and beta diversity of stream insect metacommunities at two spatial levels

The hypotheses that beta diversity should increase with decreasing latitude and increase with spatial extent of a region have rarely been tested based on a comparative analysis of multiple datasets, and no such study has focused on stream insects. We first assessed how well variability in beta diversity of stream insect metacommunities is predicted by insect group, latitude, spatial extent, altitudinal range, and dataset properties across multiple drainage basins throughout the world. Second, we assessed the relative roles of environmental and spatial factors in driving variation in assemblage composition within each drainage basin. Our analyses were based on a dataset of 95 stream insect metacommunities from 31 drainage basins distributed around the world. We used dissimilarity‐based indices to quantify beta diversity for each metacommunity and, subsequently, regressed beta diversity on insect group, latitude, spatial extent, altitudinal range, and dataset properties (e.g., number of sites and percentage of presences). Within each metacommunity, we used a combination of spatial eigenfunction analyses and partial redundancy analysis to partition variation in assemblage structure into environmental, shared, spatial, and unexplained fractions. We found that dataset properties were more important predictors of beta diversity than ecological and geographical factors across multiple drainage basins. In the within‐basin analyses, environmental and spatial variables were generally poor predictors of variation in assemblage composition. Our results revealed deviation from general biodiversity patterns because beta diversity did not show the expected decreasing trend with latitude. Our results also call for reconsideration of just how predictable stream assemblages are along ecological gradients, with implications for environmental assessment and conservation decisions. Our findings may also be applicable to other dynamic systems where predictability is low.     

Identifying resilience mechanisms to recurrent ecosystem perturbations

The complex nature of ecological systems limits the unambiguous determination of mechanisms that drive resilience to natural disturbance or anthropogenic stress. Using eight-year time series data from boreal lakes with and without bloom formation of an invasive alga (Gonyostomum semen, Raphidophyceae), we studied resilience of phytoplankton communities in relation to recurring bloom impacts. We first characterized phytoplankton community dynamics in both lake types using univariate metrics of community structure (evenness, species richness, biovolume and Simpson diversity). All metrics, except species richness, were substantially altered and showed an inherent stronger variability in bloom lakes relative to reference lakes. We assessed resilience mechanisms using a multivariate time series modelling technique. The models captured clear successional dynamics of the phytoplankton communities in all lakes, whereby different groups of species were substituted sequentially over the ice-free period. The models also identified that G. semen impacts in bloom lakes were only manifested within a single species group, not across species groups, highlighting the rapid renewal of the phytoplankton communities upon bloom collapse. These results provide empirical support of the cross-scale resilience model. Cross-scale resilience could provide an explanation for the paradox that similar species richnesses are seen in bloom-forming lakes and reference lakes despite the clear difference between the community features of the two different sets of lakes investigated.     

Detecting spatial regimes in ecosystems

Research on early warning indicators has generally focused on assessing temporal transitions with limited application of these methods to detecting spatial regimes. Traditional spatial boundary detection procedures that result in ecoregion maps are typically based on ecological potential (i.e. potential vegetation), and often fail to account for ongoing changes due to stressors such as land use change and climate change and their effects on plant and animal communities. We use Fisher information, an information theory‐based method, on both terrestrial and aquatic animal data (U.S. Breeding Bird Survey and marine zooplankton) to identify ecological boundaries, and compare our results to traditional early warning indicators, conventional ecoregion maps and multivariate analyses such as nMDS and cluster analysis. We successfully detected spatial regimes and transitions in both terrestrial and aquatic systems using Fisher information. Furthermore, Fisher information provided explicit spatial information about community change that is absent from other multivariate approaches. Our results suggest that defining spatial regimes based on animal communities may better reflect ecological reality than do traditional ecoregion maps, especially in our current era of rapid and unpredictable ecological change.     

False Correlates of Nested Patterns: A Case Study Using Temporary Pond Microcrustaceans

Nestedness analysis is a popular tool for inferring spatial species distributions, and therefore has management and conservation relevance. Ecologists frequently compute nestedness and subsequently use Spearman rank correlations for inferring relationships between the observed nested ranks of sites with biogeographic and environmental variables. Using temporary pond microcrustaceans hatched from microcosms as a case study, this paper shows that the application of this method can be problematic. While the overall degree and significance of nestedness was robust against a statistical error, the results obtained from randomly generated matrices, in which community structure from the original microcrustacean incidence matrix was maintained (fixed rows –fixed columns constraints), showed that rank correlations of observed nested patterns can be vulnerable to a Type 1 error (detecting an effect when there is none). Using expected nestedness patterns derived from rarefied original matrices to control for sample size effects did not change this result. This problem may have arisen as a result of a quantitative bias related to the disproportionate impact of rank positions of individual ponds in the analysis. Future extensive simulations studies, involving different community structures, should help identify the general reliability of rank correlation results in nestedness analyses. (© 2009 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Inferring the Relative Resilience of Alternative States

Ecological systems may occur in alternative states that differ in ecological structures, functions and processes. Resilience is the measure of disturbance an ecological system can absorb before changing states. However, how the intrinsic structures and processes of systems that characterize their states affects their resilience remains unclear. We analyzed time series of phytoplankton communities at three sites in a floodplain in central Spain to assess the dominant frequencies or “temporal scales” in community dynamics and compared the patterns between a wet and a dry alternative state. The identified frequencies and cross-scale structures are expected to arise from positive feedbacks that are thought to reinforce processes in alternative states of ecological systems and regulate emergent phenomena such as resilience. Our analyses show a higher species richness and diversity but lower evenness in the dry state. Time series modeling revealed a decrease in the importance of short-term variability in the communities, suggesting that community dynamics slowed down in the dry relative to the wet state. The number of temporal scales at which community dynamics manifested, and the explanatory power of time series models, was lower in the dry state. The higher diversity, reduced number of temporal scales and the lower explanatory power of time series models suggest that species dynamics tended to be more stochastic in the dry state. From a resilience perspective our results highlight a paradox: increasing species richness may not necessarily enhance resilience. The loss of cross-scale structure (i.e. the lower number of temporal scales) in community dynamics across sites suggests that resilience erodes during drought. Phytoplankton communities in the dry state are therefore likely less resilient than in the wet state. Our case study demonstrates the potential of time series modeling to assess attributes that mediate resilience. The approach is useful for assessing resilience of alternative states across ecological and other complex systems.     

Fire legacies in eastern ponderosa pine forests

Disturbance legacies structure communities and ecological memory, but due to increasing changes in disturbance regimes, it is becoming more difficult to characterize disturbance legacies or determine how long they persist. We sought to quantify the characteristics and persistence of material legacies (e.g., biotic residuals of disturbance) that arise from variation in fire severity in an eastern ponderosa pine forest in North America. We compared forest stand structure and understory woody plant and bird community composition and species richness across unburned, low‐, moderate‐, and high‐severity burn patches in a 27‐year‐old mixed‐severity wildfire that had received minimal post‐fire management. We identified distinct tree densities (high: 14.3 ± 7.4 trees per ha, moderate: 22.3 ± 12.6, low: 135.3 ± 57.1, unburned: 907.9 ± 246.2) and coarse woody debris cover (high: 8.5 ± 1.6% cover per 30 m transect, moderate: 4.3 ± 0.7, low: 2.3 ± 0.6, unburned: 1.0 ± 0.4) among burn severities. Understory woody plant communities differed between high‐severity patches, moderate‐ and low‐severity patches, and unburned patches (all p < 0.05). Bird communities differed between high‐ and moderate‐severity patches, low‐severity patches, and unburned patches (all p < 0.05). Bird species richness varied across burn severities: low‐severity patches had the highest (5.29 ± 1.44) and high‐severity patches had the lowest (2.87 ± 0.72). Understory woody plant richness was highest in unburned (5.93 ± 1.10) and high‐severity (5.07 ± 1.17) patches, and it was lower in moderate‐ (3.43 ± 1.17) and low‐severity (3.43 ± 1.06) patches. We show material fire legacies persisted decades after the mixed‐severity wildfire in eastern ponderosa forest, fostering distinct structures, communities, and species in burned versus unburned patches and across fire severities. At a patch scale, eastern and western ponderosa system responses to mixed‐severity fires were consistent.     

The influence of Procambarus clarkii (Cambaridae,Decapoda) on water quality and sediment characteristics in a Spanish floodplain wetland

This study describes the effects of the American red swamp crayfish, Procambarus clarkii Girard, on water quality and sediment characteristics in the Spanish floodplain wetland, Las Tablas de Daimiel National Park. Our short term enclosure study during a summer drawdown revealed that crayfish acted as a nutrient pump that transformed and translocated sediment bound nutrients to the water column. Water quality impoverishment was mainly due to the increase of dissolved inorganic nutrients (soluble reactive phosphorus and ammonia), and a significant increase of total suspended solids occurred likely as a result of crayfish associated bioturbation. At the same time, crayfish reduced the content of organic matter in the sediment and we observed a slight increase of total sediment phosphorus and nitrogen content as a result of crayfish benthic activity. P. clarkii effects, in terms of internal nutrient loading (229.91 mg TP m^–2 d^–1), were shown to be important on a local scale, indicating the significance of internal nutrient supply to water column primary producers particularly under low external supply (summer). Extrapolations to the whole ecosystem, however, revealed a negligible crayfish contribution (0.06%) to total internal nutrient loading (0.035 mg TP m^–2 d^–1). Hence, crayfish spatial heterogeneity patterns are important in global and local matter fluxes and nutrient cycles in wetlands.